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Evolution of tree killing in bark beetles (Coleoptera: Curculionidae): trade-offs between the maddening crowds and a sticky situation

Published online by Cambridge University Press:  11 June 2013

B.S. Lindgren  and
K.F. Raffa
B.S. Lindgren*
Affiliation:
Natural Resources and Environmental Studies Institute, University of Northern British Columbia, Prince George V2N 5A4, British Columbia, Canada
K.F. Raffa
Affiliation:
Department of Entomology, University of Wisconsin–Madison, Madison, Wisconsin 53706, United States of America
*
1Corresponding author (e-mail: [email protected]).
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Abstract

Bark beetles (Coleoptera: Curculionidae: Scolytinae) play important roles in temperate conifer ecosystems, and also cause substantial economic losses. Although their general life histories are relatively similar, different species vary markedly in the physiological condition of the hosts they select. Most of ∼6000 known species colonise dead or stressed trees, a resource they share with a large diversity of insects and other organisms. A small number of bark beetle species kill healthy, live trees. These few are of particular interest as they compete directly with humans for resources. We propose that tree killing evolved when intense interspecific competition in the ephemeral, scarce resource of defence-impaired trees selected for genotypes that allowed them to escape this limitation by attacking relatively healthy trees. These transitions were uncommon, and we suggest they were facilitated by (a) genetically and phenotypically flexible host selection behaviours, (b) biochemical adaptations for detoxifying a wide range of defence compounds, and (c) associations with symbionts, which together aided bark beetles in overcoming formidable constitutive and induced host defences. The ability to detoxify terpenes influenced the evolutionary course of pheromonal communication. Specifically, a mate attraction system, which was exploited by intraspecific competitors in locating poorly defended hosts, became a system of cooperative attack in which emitters benefit from the contributions responders make in overcoming defence. This functional shift in communication was driven in part by linkage of beetle semiochemistry to host defence chemistry. Behavioural and phenological adaptations also improved the beetles’ abilities to detect when tree defences are impaired, and, where compatible with life history adaptations to other selective forces, for flight to coincide with seasonally predictable host stress agents. We propose a conceptual model, whereby the above mechanisms enable beetles to concentrate on those trees that offer an optimal trade-off between host defence and interspecific competition, along dynamic gradients of tree vigour and stand-level beetle density. We offer suggestions for future research on testing elements of this model.

Résumé

Les scolytes (Coleoptera: Curculionidae: Scolytinae) tiennent des rôles importants dans les écosystèmes tempérés de conifères et y causent aussi de sérieuses pertes économiques. Bien que leurs cycles biologiques soient généralement assez semblables, les différentes espèces diffèrent considérablement par les conditions physiologiques des hôtes qu'elles choisissent. La plupart des quelque 6000 espèces connues colonisent les arbres morts ou soumis à des stress, une ressource qu'ils partagent avec une grande variété d'insectes et d'autres organismes. Un petit nombre d'espèces de scolytes tuent des arbres vivants et sains. Ces dernières sont d'intérêt particulier car elles font compétition directement avec les humains pour les ressources. Nous pensons que la stratégie de tuer les arbres s'est développée lorsqu'une intense compétition interspécifique pour la ressource éphémère et rare d'arbres aux défenses affaiblies a favorisé la sélection de génotypes qui permettaient d’échapper à ces restrictions en attaquant des arbres relativement sains. Ces transitions se sont produites rarement et nous croyons qu'elles ont été facilitées par a) des comportements de sélection des hôtes génétiquement et phénotypiquement flexibles, b) des adaptations biochimiques pour la détoxification d'une gamme étendue de composés de défense et c) des associations avec des symbiontes qui ensemble ont aidé les scolytes à surmonter les formidables défenses constitutives et induites de l'hôte. La capacité de détoxifier les terpènes a influencé le cours de l’évolution de la communication par phéromones. De manière plus spécifique, un système d'attraction du partenaire qui a été exploité par des insectes en compétition intraspécifique pour localiser les hôtes à défenses affaiblies est devenu un système d'attaque coopérative dans lequel ceux qui émettent bénéficient des contributions faites par ceux qui répondent pour ainsi surmonter les défenses. Ce déplacement fonctionnel dans la communication s'est opéré en partie par le lien établi entre la sémiochimie du coléoptère et la chimie de défense de son hôte. Des adaptations comportementales et phénologiques ont aussi amélioré la capacité des coléoptères à discerner quand les défenses de l'arbre sont affaiblies et de faire coïncider leur vol avec les agents de stress de l'hôte prévisibles au cours de la saison, lorsque cela est compatible avec les adaptations du cycle biologique aux autres forces de sélection. Nous proposons un modèle conceptuel dans lequel les mécanismes décrits ci-haut permettent aux coléoptères de se concentrer sur les arbres qui offrent un compromis optimal entre la défense de l'hôte et la compétition interspécifique, le long de gradients dynamiques de vigueur des arbres et de densité des coléoptères dans le peuplement. Nous présentons des suggestions pour des recherches ultérieures pour tester les éléments de ce modèle.

Type
Review
Information
The Canadian Entomologist , Volume 145 , Issue 5 , October 2013 , pp. 471 - 495
Copyright
Copyright © Entomological Society of Canada 2013 

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